Low effective mechanical advantage of giraffes' limbs during walking reveals trade-off between limb length and locomotor performance

Giraffes (Giraffa camelopardalis) possess specialized locomotor morphology, namely elongate and gracile distal limbs. While this contributes to their overall height and enhances feeding behavior, we propose that the combination of long limb segments and modest muscle lever arms results in low effective mechanical advantage (EMA, the ratio of in-lever to out-lever moment arms), when compared with other cursorial mammals To test this, we used a combination of experimentally measured kinematics and ground reaction forces (GRFs), musculoskeletal modeling, and inverse dynamics to calculate giraffe forelimb EMA during walking. Giraffes walk with an EMA of 0.34 (+/- 0.05 SD), with no evident association with speed within their walking gait. Giraffe EMA was about four times lower than expectations extrapolated from other mammals, ranging from 0.03 to 297 kg, and this provides further evidence that EMA plateaus or even diminishes in mammals exceeding horse size. We further tested the idea that limb segment length is a factor which determines EMA, by modeling the GRF and muscle moment arms in the extinct giraffid Sivatherium giganteum and the other extant giraffid, Okapia johnstoni. Giraffa and Okapia shared similar EMA, despite a four to sixfold difference in body mass (Okapia EMA = 0.38). In contrast, Sivatherium, sharing a similar body mass with Giraffa, had greater EMA (0.59), which we propose reflects behavioral differences, such as a somewhat increased capability for athletic performance. Our modeling approach suggests that limb length is a determinant of GRF moment arm magnitude and that unless muscle moment arms scale isometrically with limb length, tall mammals are prone to low EMA.